Diptera: Sarcophagidae
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The JapanSocietyJapan Society ofMedicalof Medical Entomology and Zoology [Med.Entomol.Zool,Nb].53 SuppL2 p,259-273 2002) Nomenclature ofMusca carnan'a Linnaeus, 1758 (Diptera: Sarcophagidae) Thomas PApE Department ofEntomology S4,edish Mitseum ofNbtural History RO. Box 5000Z - SE 104 05 Stockholm, Sweden E-mail:thomas,[email protected] (Received: 7 November 2001; Accepted: 1O December 200l) Key words: nomenclature, Diptera, Sarcophagidae, priority; stability Abstract: The nomenelature of lhsca earnaria Linnaeus, 1758, currently in the SZireophaga genus Meigen, 1826, is suceinctly reviewed. The original type material is discussedand argued to consist ofat one North least American as well as several European specimens, with most ofthe latter no longer in existence. The lectotype designation and the resulting nomenclatural changes are discussedin light of recent critique, The lectotype designation is in agreement with the relevant articles of the International Code of Zoological abandoning the Nomenclature, but by (at that time) current usage it may arguably be considered too rigid an application of the Principle of Priority. As such, the lectotype designation may not have been optimal when proposed, yet the general acceptance in the scientific Iiterature emerging during the Iast fifteen years provides hope fbr nomenclatural stability, and no contrary action is recommended. INTRODUCTION Linnaean types still have the capacity to stir up nomenclature. Some of these old types have remained unrevised, probably from a combination of awe and tradition, while others have been revised only recently (e.g., Day and Fitton, 1978; Pont, 1981; Thempson et aL 1982; Richet, 1987). The nomenclature of the only nominal flesh fly taxon proposed by Linnaeus: Musca carnaria Linnaeus, 1758, was revised by Richet (1987), who designated a lectotype chosen among two specimens in the Linnaean collection. Lehrer (2000) criticised this designation, as he found the nomenclatural changes unacceptably at odds with current usage. Neither author, however, has fully acknowledged that zoological nomenclature is as much a study ofhistory as oftaxonomy and biology, and it is the purpose of the present paper to review succinctly the nomenclature of the nominal taxon Musca carnaria Linnaeus, 1758 (currently in genus Sbrcophcrga Meigen) and to discuss Richet's lectotype designation in the light of the critique addressed by Lehrer, In so doing, I refer to IUchet (1987) and Lehrer (2000) without redundantly repeating the year, and I cite the relevant Articles of the International Code of Zoological Nomenclature (ICZN) as they appear in the 4th edition (ICZN 1999), with the equivalent and mostly identical Articles of the 3rd edition (ICZN NII-Electronic Library Service The JapanSocietyJapan Society ofMedicalof Medical Entomology and Zoology 260 Med. EntomoL. Zool. 1985), which were in effect when Richet published his lectotype designation, in brackets, TYPE MATERIAL Linnaeus (1758: 596) placed Musca carnaria as the 53rd species ofthe genus Musca in the 10th edition of his unsurpassed Systema Nbturae. Following the description and list of "Hbbitat references he gave the information: in Cadaveribus Europae, etiam Amerieae. Ktilm." In other words, Linnaeus considered this fly to be a carrion breeder distributed in Europe as well as in [North] Arnerica. Linnaeus gave as a source of (part of) this information "Kalm", which refers to Per Kalm, a Swedish naturalist and Linnaean disciple, who collected in North America and published on the North American biota including its insect fauna (Kerklconen, 1959; for several additional references see Evenhuis, 1997). The brief mention of a carrion-breeding habit ("Hbbitat in Cadaveribus", note the slight difference from the corresponding entry in Fauna Svecica, where Linnaeus (1746) gave "Habitat in carnibus recentibus") possibly originated from such infbrmation in the literature sources cited by Linnaeus; yet it may as well refer to original observations of Iarvae in decaying meat, made by Linnaeus himselC or more probably by De Geer, a contemporary naturalist and fe11ow countryman to Linnaeus and a remarkably keen observer of natural history (Persson et aL, 1984), De Geer (1776) provided (unavailable) names fbr what he considered two species of viviparous flies, Musca vivipara mojor, a replacement name given by De Geer fbr Miisca carnaria Linnaeus, and Miisca vivipara minor. De Geer gave detailed illustrations and descriptions undoubtedly referable to larvae of SZircophaga sp., here fbllowing the broad definition of this genus outlined by Pape (1996). The Swedish material studied by De Geer could belong to one or more species from several subgenera, including Sarcophaga (sensu strieto), which by default of Partington's (1837) typification includes S carnaria. De Geer's detailed plates even show the first illustration ever made of male sarcophagid genitalia, which I have identified as belonging to the species Stircophaga (Pandelleisca) similis Meade (Pape, 1993). Linnaeus had close ties to De Geer and it is likely that they shared information prior to the publication of the 10th edition of Systema AJOturae. In a nomenclatural context, however, Linnaeus' biological note is a poor pointer to real spe ¢ imens and can hardly be considered unambiguous evidence fbr an extensive type senes. The mention of [North] America as part ofthe distribution would suggest a whole range of sarcophagine subgenera, though not Sbrcophaga (sensu stricto), which only recently was introduced to eastern Canada (Pape, 1996, see note on S subvicina RohdendorC p. 14). Interestingly, we have a clue to pinpoint what appears to be ari American contingent to the type series. The De Geer collection at the Swedish Museum ofNatural History contains one female specimen standing under the name Musca vivipara mojor and carrying a handwritten, "Carriaria.". irregular label (handstyle not interpreted) giving I have earlier left this female NII-Electronic Library Service The JapanSocietyJapan Society ofMedicalof Medical Entomology and Zoology 261 Vbl,53 Suppl,2, 2002 unidentified 1993), (Pape, but a re-examination of the specimen has convinced me that it belongsto Sarcophaga (?Veobellieria) bullata Parker, which is a common species in eastern Nonh America breeding in garbage and carrion (specimen compared with females from a bred series from South Carolina). De Geer (1776) stated Pennsylvania to be part of the distribution of his Musca vivipara mojor, and it is not unlikely that the female was collected by the Linnaean disciple Per Kalm. Linnaeus is known to have described insect species on specimens m De Geer's collection, and he might have Iearned about or even studied the femaleSbrcophaga bullata. In any case, De Geer (1776) was soon added as an authoritative reference to Musca carnaria, and the American-European distribution was maintained (e.g., Gmelin, 1790). The literaturesources Frisch, (Aldrovandus, Jonston, Linnaeus, Reaumur, Roesslin) cited by Linnaeus (1758) are explicit references to earlier treatments of what Linnaeus considered to be Sbrcophaga carnaria, and as such the specimens treated (described, illustrated) therein, irrespectiye of their historical fate, are to be regarded as part of the type series fbllowing Article 72.4.1 [72(b)(i)]. Reaumur (1738), fbr example, extensively described what he called "des "des mouches vivipares" and mouches grises" (pp. 408-430), and his illustrations of females with distinct black thoracic stripes (pl. 29, figs 4-6) and his carefu1 description of recessed posterior larval spiracles (p. 428) accompanied with an illustration showing the almost vertical spiracular slits (pl. 29, fig. 2) leave no doubt that he was studying one or more species of Sbrcophaga (sensu lato). Reaumur caught gravid females indoors searching for to larviposit places (p. 408), which most likely included Sbreophaga (Liop)tgia) argyrostoma (Robineau-Desvoidy), which is one of the few European species of Sarcophagidae that frequents 'meat' houses to reproduce in mammal carrion, i.e., (Leclercq and Verstraeten, 1988; Schumann, 1990). No attempt has been made here to trace fUrther these pre-Linnaean concepts, but Linnaeus evidently based his concept ofMusca carnaria not only on material in his own collection but, through his literature sources, on an unlrnown number of additional specimens that are likely to represent additional species ofSbrcophaga (s.L). LECTOTYPEDESIGNAT-ION Haliday (1851) was the first to study and publish explicitly on the Linnaean material of Musca carnaria housed at the Linnean Society in London. He gives only the brief statement ""53. = Carnaria," Sarcophaga id. Mg.", which does not qualify as a lectotype designation and in itself is hardly infbrmative considering the insuffTicient knowledge available for Skercophaga carnaria sensu Meigen (and other 19th century authors) as is elaborated below. Tbwnsend "Musca (1 938: 64), under his entry fbr Sarcophaga, gives carnaria Linne [. , .] Ht - Origin, Sweden; location, Uppsala or lost". While this is an explicit reference to a holotype ("Ht"), it appears from the context that it is coniectural, and it does not refer to one particular specimen, for which reasons it does not fu1fi1 the requirements for a pre-2000 lectotype NII-Electronic Library Service The JapanSocletyJapan Society ofMedicalof Medical Entomology and Zoology 262 Med. Entomol. Zool. designation. When Richet (1987) studied the Linnaean material ofM carnaria more than a century after Haliday (brought to him during a visit at